Physiology of Smooth Muscle

Structure of smooth muscle

composed of spindle-shaped fibers

diameter of 2-10 µm and lengths of several hundred µm

lack the coarse CT sheaths of skeletal muscle, but have fine endomysium

are typically organized into two layers (“longitudinal” and “circular”) of closely apposed fibers

found in walls of hollow organs (except the heart)

have essentially the same contractile mechanisms as skeletal muscle

microscopic anatomy

SR is less developed than in skeletal muscle and lacks a specific pattern

T tubules are absent

plasma membranes have pouchlike infoldings called caveoli

Ca2+ is sequestered in the extracellular space near the caveoli, allowing rapid influx when channels are opened

no visible striations and no sarcomeres

thin and thick filaments are present

organization of myofilaments

ratio of thick to thin filaments is much lower than in skeletal muscle

thick filaments have heads along their entire length

there is no troponin complex

thick and thin filaments are arranged diagonally, causing smooth muscle to contract in a corkscrew manner

noncontractile intermediate filament bundles attach to dense bodies (analogous to Z discs; composed of α-actinin) at regular intervals

Innervation

no neuromuscular junctions

innervating nerves have bulbous swellings called varicosities

varicosities release neurotransmitters into wide synaptic clefts called diffuse junctions

Contraction

visceral contractions

when the longitudinal layer contracts, the organ dilates and contracts

when the circular layer contracts, the organ elongates

peristalsis

alternating contractions and relaxations of smooth muscles that mix and squeeze substances through the lumen of hollow organs

whole sheets of smooth muscle exhibit slow, synchronized contraction

contract in unison, reflecting their electrical coupling with gap junctions

action potentials are transmitted from cell to cell

some smooth muscle cells:

act as pacemakers and set the contractile rate for whole sheets of muscle

are self-excitatory and depolarize without external stimuli

contractile mechanism

smooth muscle contractile mechanism

actin and myosin interact according to the sliding filament mechanism

final trigger for contractions is a rise in intracellular Ca2+

Ca2+ is released from the SR and from the extracellular space

Ca2+ interacts with calmodulin and myosin light chain kinase to activate myosin

role of Ca2+

binds to calmodulin and activates it

activated calmodulin activates the kinase enzyme

activated kinase transfers phosphate from ATP to myosin cross bridges

phosphorylated cross bridges interact with actin to produce shortening

muscle relaxes when intracellular Ca2+ levels drop

signal transductions mechanisms

smooth muscle signal transduction pathways

phosphatidylinositol pathway

GS-protein-coupled pathway

nitric oxide (NO)-cGMP pathway

special features of SM contraction

unique characteristics:

smooth muscle tone

slow, prolonged contractile activity

low energy requirements

response to stretch

smooth muscle exhibits a phenomenon called stress-relaxation response in which:

responds to stretch only briefly, and then adapts to its new length

new length, however, retains its ability to contract

this enables organs such as the stomach and bladder to temporarily store contents

Hyperplasia

certain smooth muscles can divide and increase their numbers by undergoing hyperplasia

shown by estrogen’s effect on the uterus

at puberty, estrogen stimulates the synthesis of more smooth muscle, causing the uterus to grow to adult size

during pregnancy, estrogen stimulates uterine growth to accommodate the increasing size of the growing fetus

Types of Smooth Muscle

Single Unit

commonly found as visceral muscle, the cells of which:

contract rhythmically as a unit

are electrically coupled to one another via gap junctions

often exhibit spontaneous action potentials

are arranged in opposing sheets and exhibit stress-relaxation response

Multiunit

located in:

large airways to the lungs

large arteries

arrector pili muscles attached to hair follicles

internal eye muscles (sphincter & dilator pupillae mm)

characteristics:

rare gap junctions

infrequent spontaneous depolarizations

structurally independent muscle fibers

rich nerve supply, which, with a number of muscle fibers, forms motor units

graded contractions in response to neural stimuli





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